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Brief history of photosynthetic organisms on earth Division: Cyanobacteria 3.45 bya = Cyanobacteria appear and introduce photosynthesis blue-green algae 1.5 bya = first Eukaryotes appeared (nuclear envelope and ER thought to come from invagination of plasma membrane) 0.9 bya = first multicellular algae (Rhodophyta - Red algae) 800 mya = earliest Chlorophyta (Green algae) 400-500 mya = plants on land – derived from Charophyceae 250 mya = earliest Heterokontophyta (Brown algae) 100 mya = earliest seagrasses (angiosperms) • Single class: Cyanophyceae • ~150 genera • ~4037 species 1 Groups (Kingdom) DOMAIN 1.Bacteria- cyanobacteria (blue green algae) 2.Archae 3.Eukaryotes 1. Alveolates- dinoflagellates 2 Domain Bacteria- unicellular, lacking a nucleus, ribosomes, tRNA, peptoglycan in the cell wall Phylum Low-GC Gram-positive- staph infections & antrax Phylum High GC- Gram-positive- tuberculosis & antibiotics Phylum Hyperthermophilic bacteria- live at high temperatures Phylum Hadobacteria-live at high temps, consume nuclear waste Phylum Cyanobacteria- blue green algae, origin of the chloroplast Phylum Spirochetes- syphilis & lime disease Phylum Chlamydias- intercellular parasites Phylum Proteobacteria- pathogens, nitrogen fixers, autotrophs, origin of mitochondria 2. Stramenopiles- diatoms, heterokonyophyta 3. Rhizaria- unicellular amoeboids 4. Excavates- unicellular flagellates 5. Plantae- rhodophyta, chlorophyta, seagrasses 6. Amoebozoans- slimemolds 7. Fungi- heterotrophs with extracellular digestion 8. Choanoflagellates- unicellular 9. Animals- multicellular heterotrophs 3 4 1 Cyanobacteria Cellular Structure: Cyanobacteria are important because: •Nucleus? • Cyanobacteria are modern representatives of the very first photosynthetic organisms on Earth. •Chloroplasts? •First organisms to have 2 photosystems and to produce organic material and to give off O 2 as a biproduct. • Pigments? •Instrumental in transforming early earths atmosphere to an oxidizing one the oxygen revolution • Carbon storage? • Most common algal group in terrestrial systems and symbiotic relationships • Flagella? 5 Cyanobacteria Cellular Structure 6 Cellular Structure: microscopic No complex organelles • Unicells in mucus envelope Thyllakoids- photosystems I & II Carboxisomes- bacterial microcompartments that contain enzymes involved in carbon fixation - concentrate CO2 for RuBisCo • Colonies • Filaments, Branched filaments most complicated form 7 8 2 Cyanophyta Cellular Structure vegetative cells- photsynthesize “facultative chemoautotropes” spores- resting stages •Many spp have the ability to photosynthesize under both aerobic and anaerobic conditions heterocysts- specialized cells for fixing nitrogen Difference is where the electrons come from: Aerobic conditions? Electron donor is Water O2 produced C0 2 + H2 0 CH2 O + O2 Anaerobic conditions? Electron donor is Hydrogen sulfide Water is produced 2H2 S + CO2 CH2 O + 2S + H2 O 9 Morphology of Cyanophyta • Cell wall made of peptidoglycan, polymer consisting of sugars and amino acids (not cellulose), similar to gram-negative bacteria • Trichome- Row of cells • Filament- Trichome within a mucilaginous sheath 10 Mucilaginous Sheath – –motion (gliding) -protection against desiccation - protection against UV irradiance Sheath is often colored: Red = acidic Blue = basic Yellow/Brown = high salt • Mucilaginous Sheath- layer of mucilage outside of cell wall • Possible to have > 1 trichome within a filament 11 12 3 False branching = True branching = outgrowth of filaments adjacent to dead or specialized cells; filament curves outgrowth from cells that change their axis of division, 90 degrees from axis of trichome Movement 1. ”Gliding” against a solid substrate -no change in shape or obvious pushing; no ability to steer, sometimes trichome rotates within sheath, and sheath is left behind as the trichome moves forward 2. Jet propulsion = excrete mucus 3. Helical species (e.g. Spirulina), use waves of contraction 4. Swimming? No idea how they do this, but evidence of chemotaxis and phototaxis. 5. Changing buoyancy 13 Buoyancy: 14 Reproduction: Cells contain gas vesicles or gas vacuole = hollow cylinders made of protein •low light? Decreased Ps metabolism of polysaccharides increase in gas vacuoles float upward •high light? Increased Ps Accumulation of polysaccharides cell pressure increases gas vacuoles shrink sink No sexual reproduction some DNA transfer has been observed, not conjugation per se. Asexual reproduction through: 1-Binary fission – dividing in two 2-Fragmentation of colonies 3-Endospores/exospores 4-Hormogonia 5-Akinetes 15 16 4 Reproduction Asexual Reproduction Hormogonia – short piece of trichome found in filaments. It detaches from parent filament and glides away 4. Hormogonia = short piece of trichome that detaches from parent filament and glides away Separation disk or “Necridium” = funky dead cell where detachment occurs 17 18 Hormogonia Reproduction Akinete 5. Akinetes = thick-walled resting spores. Packed with energy reserves; dense = sink to the bottom when released. Triggered by unfavorable conditions, can remain viable for years. H 19 A - akinete 20 5 Earliest evidence of Cyanobacteria comes from stromatolites: Precambrian- 3.5 BYA Wide Range of Habitats •Freshwater lakes •Terrestrial soils hotsprings at Yellowstone Park • Marine systems (intertidal, open ocean) • Extreme environments (e.g. salt flats, hot springs, glaciers, etc.) • Endosymbiotic: diatoms, sponges, tunicates, lichens, polar bear fur, bryophytes, gymnosperms, angiosperms 21 Stromatolites • Stromatolites are produced by successive deposition through “grain trapping” or calcification: • Mucilaginous sheath of cyanos physically blocks the movement coarse gain sediments and laminates it to the surface of the stromatolite • Attract and bind Ca ions to negatively charged sites • Locations : hypersaline seas (Shark Bay,western Aus.), frozen lakes (Antarctic), hot springs (Yellowstone), the Caribbean •Most cyano’s active during the day= layers count the numbers of days •Also they grow up toward the sun and are directional toward the sun (=can document annual motion of sun) 23 • Layered calcareous mounds that contain fossils of cyanobacteria that look like Oscillatoria 22 Another big reason Cyanobacteria are cool and ecologically important: Nitrogen fixation Many are able to fix nitrogen = convert atmospheric N2 (N N) to a usable form (Ammonium: NH 4 +) using enzyme nitrogenase N can be limiting; necessary for the production of amino acids Only cyanobacteria and proteobacteria can fix N; BUT cyanobacteria also produce O2 during photosynthesis This is a trick, because O2 inactivates nitrogenase How do they do it??? 24 6 1. Spatial separation of functions: 2. Temporal separation of functions: Heterocyst = special cells for N fixation. • • Thick-walled, and larger than other vegetative cells; hollow looking Fix N in the dark, Ps in the daytime; Every N-fixing cyanobacteria fits into these two categories except: Trichodesmium = marine, colonial species; fix nitrogen under aerobic conditions in the light through division of • Not capable of dividing labor among cells within a filament (no heterocysts)! • Not photosynthetic, so no CO2 fixation or O2 production • Microplasmedesmat a connect to other cells in filament 25 26 Domain Bacteria- unicellular, lacking a nucleus, ribosomes, tRNA, peptoglycan in the cell wall The dark side of cyanobacteria? Phylum Cyanobacteria- 4037 species Class Cyanophyceae- 4037 species, 7 orders Cyanobacterial blooms = death and destruction Swimmer’s itch = Lyngbia releases chemicals Order Chroococcales- 954 species filamentous, no specialized structures Cyanotoxins: released by animal ingestion neurotoxins (e.g. Anabaena, Oscillatoria ) and hepatotoxins (e.g. Microcystis, Nostoc ) (death to mammals, birds, fishes , no known human deaths) Order Oscillatoriales –926 species filamentous, no specialized structures Order Nostocales- 1,297 species filamentous, specialized structures, may have true branching Oscillatoria 27 28 7 Order Chroococcales Genus Spirulina Domain Bacteria- unicellular, lacking a nucleus, ribosomes, tRNA, peptoglycan in the cell wall • Filament is spiral •Major food source for flamingos • Commercial food source • No heterocysts or akinetes Phylum Cyanobacteria- blue green algae, origin of the chloroplast • benthic or plantonic, freshwater, thermal & mineral springs, coastal & brackish waters • 52 species Class CyanophyceaeOrder Chroococcales- filamentous, no specialized structures Order Oscillatoriales -filamentous, no specialized structures Order Nostocales- filamentous, specialized structures, may have true branching 29 Order Oscillatoriales Order Oscillatoriales Genus Oscillatoria 30 - Unbranched filaments -No readily identifiable sheath, but is present -No specialized cells -Fix nitrogen -Early precambrian fossil stromatolites -Planktonic, form mats on mud, stone, sand, in freshwater, brachish, marine -137 species 31 GenusTrichodesmium - Important marine N fixer - Segregates N fixing within colonies (lacks heterocysts) - More Phycoerythrin than Phycocyanin Red color, gave the Red Sea its name - Strictly planktonic- marine & fresh water -10 species 32 8 Domain Bacteria- unicellular, lacking a nucleus, ribosomes, tRNA, peptoglycan in the cell wall Phylum Cyanobacteria- blue green algae, origin of the chloroplast Class Cyanophyceae- Order Nostocales Genus Nostoc Order Chroococcales- filamentous, no specialized structures • Fixes nitrogen • Has heterocysts in the middle of filaments •Akinates midway between heterocysts •94 species •Freshwater Order Oscillatoriales -filamentous, no specialized structures Order Nostocales- filamentous, specialized structures, may have true branching 33 Order Nostocales Genus Anabaena 34 Order Nostocales • Produces neurotoxins that become release when they are ingested by animals • Fixes nitrogen • Has heterocysts in the middle of filaments • Akinates adjacent to heterocysts • Planktonic, or periphytic, benthic, soils, • 117 species 35 Genus Calothrix • Fixes nitrogen • Has heterocysts, often at the base of filaments • Filaments taper at ends • Periphytic on algae, aquatic plants, stone, wood, high intertidal •132 species 36 9 Order Nostocales Cylindrospermum Order Nostocales • Produces neurotoxins that are released when they are ingested. • Fixes nitrogen • Has heterocyst at the base of large akinetes •Periphytic or soils •49 species 37 GenusTolypothrix • Fixes nitrogen • Has heterocysts • Exhibits false branching • 60 species •Mainly submerged, also aerophytic habitats 38 Order Nostocales Stigonema - Has heterocysts - Exhibits True Branching - Often found as phycobiont in lichens, terrestrial & soil, freshwater - 62 species 39 10